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Estudio Del Sistema Del Interferón En Células Aviares Infectadas Con El Reovirus Aviar S1133

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Estudio Del Sistema Del Interferón En Células Aviares Infectadas Con El Reovirus Aviar S1133 TESIS DOCTORAL ESTUDIO DEL SISTEMA DEL INTERFERÓN EN CÉLULAS AVIARES INFECTADAS CON EL REOVIRUS AVIAR S1133 Irene Lostalé Seijo DEPARTAMENTO DE BIOQUÍMICA E BIOLOXÍA MOLECULAR CENTRO SINGULAR DE INVESTIGACIÓN EN QUÍMICA BIOLÓXICA E MATERIAIS MOLECULARES SANTIAGO DE COMPOSTELA 2015 TESIS DOCTORAL ESTUDIO DEL SISTEMA DEL INTERFERÓN EN CÉLULAS AVIARES INFECTADAS CON EL REOVIRUS AVIAR S1133 Fdo.: Irene Lostalé Seijo DEPARTAMENTO DE BIOQUÍMICA E BIOLOXÍA MOLECULAR CENTRO SINGULAR DE INVESTIGACIÓN EN QUÍMICA BIOLÓXICA E MATERIAIS MOLECULARES SANTIAGO DE COMPOSTELA 2015 El Dr. Francisco Javier Benavente Martínez, Catedrático del Departamento de Bioquímica y Biología Molecular de la Universidad de Santiago de Compostela, INFORMA: Que el trabajo de Tesis Doctoral titulado “Estudio del sistema del interferón en células aviares infectadas con el reovirus aviar S1133” que presenta Irene Lostalé Seijo para optar al grado de Doctor en Bioquímica y Biología Molecular, ha sido realizado en este Departamento bajo mi dirección y la del Dr. José Manuel Martínez Costas y, considerando que se haya concluido, autorizo su presentación para que pueda ser evaluada por el Tribunal correspondiente. Y para que así conste, firmo el presente informe: Fdo.: Francisco Javier Benavente Martínez Fdo.: José Manuel Martínez Costas Fdo.: Irene Lostalé Seijo Santiago de Compostela, 23 de Febrero de 2015. El trabajo recogido en la presente memoria ha sido realizado en el Departamento de Bioquímica y Biología Molecular de la Universidad de Santiago de Compostela, bajo la dirección de los doctores Francisco Javier Benavente Martínez y José Manuel Martínez Costas. Irene Lostalé Seijo ha disfrutado de una beca para la realización de estudios de Tercer Ciclo de la Xunta de Galicia (2006), una beca para la Formación de Profesorado Universitario del Ministerio de Educación y Ciencia (AP2006-03859) y dos becas de investigación de la Diputación Provincial de A Coruña (convocatorias 2011 y 2013). ESTUDIO DEL SISTEMA DEL INTERFERÓN EN CÉLULAS AVIARES INFECTADAS CON EL REOVIRUS AVIAR S1133 RESUMO Os reovirus aviarios son virus sen envoltura lipídica e xenoma de ARN bicatenario que infectan a aves, causando enfermidades coma a artrite infecciosa ou a síndrome de malabsorción. Neste traballo estudouse a resposta inmune innata que este virus desencadea en dous tipos celulares do seu hospedador natural, o polo, e comparouse coa xerada por outros virus coma o virus vaccinia e o virus da estomatite vesicular. O reovirus aviario é o único dos virus estudados capaz de inducir a produción de interferón (IFN) e aumentar os niveis da proteína quinasa R (PKR) en cultivos primarios de fibroblastos embrionarios de polo (CEF), mentres que é incapaz de facelo na liña celular DF1 derivada destes, o que podería depender de diferenzas nos niveis dos receptores de recoñecemento de patrón (PRRs) ou a unha menor eficiencia de infección destas células. A indución de IFN en CEF infectadas co reovirus aviario depende da decapsidación do virus pero non da expresión dos seus xenes. O reovirus aviario é resistente ao IFN e é capaz de impedir a activación de PKR e a fosforilación de eIF2, ao contrario que os outros dous virus estudados. Esta capacidade para impedir a activación de PKR depende da capacidade da proteína A para unir ARN bicatenario. Por outra banda, observouse que o pretratamento con IFN de células DF1 aumenta a cantidade de células infectadas co reovirus aviario mediante un mecanismo independente da activación das caspasas. Finalmente, estudáronse algúns posibles mecanismos da inhibición da síntese das proteínas celulares durante a infección con reovirus aviario. PALABRAS CLAVE virus, inmunidade innata, reovirus aviario, interferón, proteína quinasa R RESUMEN Los reovirus aviares son virus desnudos con genoma de ARN bicatenario que infectan a aves, causando enfermedades como la artritis infecciosa o el síndrome de malabsorción. En este trabajo se ha estudiado la respuesta inmune innata que este virus desencadena en dos tipos celulares de su hospedador natural, el pollo, y se ha comparado con la que generan otros virus como el virus vaccinia y el virus de la estomatitis vesicular. El reovirus aviar es el único de los virus estudiados capaz de inducir la producción de interferón (IFN) y aumentar los niveles de la proteína quinasa R (PKR) en cultivos primarios de fibroblastos embrionarios de pollo (CEF), mientras que es incapaz de hacerlo en la línea celular DF1 derivada de éstos, lo que podría depender de diferencias en los niveles de los receptores de reconocimiento de patrón (PRRs) o a una menor eficiencia de infección de estas células. La inducción de IFN en CEF infectadas con reovirus aviar depende de la decapsidación del virus pero no de la expresión de sus genes. El reovirus aviar es resistente al IFN y es capaz de impedir la activación de PKR y la fosforilación de eIF2, al contrario que los otros dos virus estudiados. Esta capacidad para impedir la activación de PKR depende de la capacidad de la proteína A para unir ARN bicatenario. Por otra parte, se ha observado que el pretratamiento con IFN de células DF1 aumenta la cantidad de células infectadas con el reovirus aviar mediante un mecanismo independiente de la activación de las caspasas. Finalmente, se han estudiado algunos posibles mecanismos de la inhibición de la síntesis de proteínas celulares durante la infección con reovirus aviar. PALABRAS CLAVE virus, inmunidad innata, reovirus aviar, interferón, proteína quinasa R 9 ABSTRACT Avian reoviruses are nonenveloped viruses containing a dsRNA genome that infect birds, causing diseases like viral arthritis or malabsorption syndrome. In this report, the innate immune response triggered by this virus on two chicken cell lines has been studied and compared to the ones caused by vaccinia virus and vesicular stomatitis virus. Only avian reovirus is able to induce interferon (IFN) secretion and protein kinase R (PKR) induction in primary chicken embryo fibroblasts (CEF), while it is unable to do so in DF1 cells, probably due to differences on pattern recognition receptors (PRRs) levels or infection efficiency. IFN induction in avian reovirus infected cells depends on viral uncoating but not on viral gene expression. Avian reovirus is resistant to IFN effects and blocks PKR activation and eIF2 phosphorylation, unlike the other viruses used on this study. The ability to inhibit PKR activation relies on A protein’s ability to sequester dsRNA. On the other hand, IFN pretreatment of DF1 cells increases the number of avian reovirus infected cells through a mechanism independent of caspase activation. Finally, several possible mechanisms of host protein shutoff during avian reovirus infection are explored. KEYWORDS virus, innate immunity, avian reovirus, interferon, protein kinase R 10 ÍNDICE ABREVIATURAS ......................................................................................................... 15 INTRODUCCIÓN ......................................................................................................... 21 1.- LA FAMILIA REOVIRIDAE ............................................................................... 23 2.- ESTRUCTURA DEL REOVIRUS AVIAR ......................................................... 24 2.1.- El genoma del reovirus aviar ....................................................................... 24 2.2.- Proteínas del reovirus aviar ......................................................................... 25 2.3.- Estructura de la partícula viral ..................................................................... 32 3.- EL CICLO REPLICATIVO DEL REOVIRUS AVIAR ....................................... 32 3.1.- Entrada y decapsidación .............................................................................. 32 3.2.- Expresión génica ......................................................................................... 34 3.3.- Morfogénesis y salida .................................................................................. 35 4.- EL SISTEMA DEL INTERFERÓN Y EL ESTADO ANTIVIRAL .................... 36 4.1.- El sistema del interferón y los interferones ................................................. 36 4.2.- Detección de la infección viral .................................................................... 38 4.3.- Señalización por interferones y creación y activación del estado antiviral . 40 5.- REOVIRUS Y RESPUESTA CELULAR ............................................................ 45 5.1.- Reovirus de mamífero ................................................................................. 45 5.2.- Reovirus aviar .............................................................................................. 46 6.- VIRUS VACCINIA Y VIRUS DE LA ESTOMATITIS VESICULAR .............. 47 6.1.- Virus vaccinia .............................................................................................. 47 6.2.- Virus de la estomatitis vesicular .................................................................. 48 OBJETIVOS .................................................................................................................. 51 MATERIALES Y MÉTODOS ..................................................................................... 55 1.- MATERIAL BIOLÓGICO ................................................................................... 57 1.1.- Células eucariotas ........................................................................................ 57 1.2.- Virus ...........................................................................................................
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